142 B. Krystufeketal. Zusammenfassung Beschreibung,Taxonomie und Verbreitungvon Talpa davidiana Wegen großer morphologischerÄhnlichkeit derSchädeldes Typus von Talpastreeti mitScaptochirus davidianus wird auf Konspezifität der beiden Taxa geschlossen. Das Becken von Talpa davidiana ist caecoidal, so daß eine Verwandtschaft mit Scaptochirus moschatus aus China ausgeschlossen wer- den kann. T davidiana wurde aus dem Grenzgebiet von Syrien und Kleinasien beschrieben und ist bisher von drei Fundorten (Umgebung von Meydanekbez, Hakkari und Tatvan) bekannt. Bei Tatvan lebt die Art sympatrisch mit Talpa levantis. Individuen von T. davidiana aus Iran und Hakkari haben eine Gesamtzahl von 44Zähnen. Als Folge des Fehlens der unteren Incisivi und der stiftförmiger Prämolaren ist die Zahnzahl bei drei Maulwürfen aus Tatvan auf 38 und beim Typus von 5. davidianus auf 39 (Asymmetrie) reduziert. References Capolongo,D.; Panasci,R. (1978): Richerche Felten,H.; Spitzenberger,F.; Storch,G. (1973): sulle populazioni di talpe dell'Italia setten- ZurKleinsäugerfaunaWest-Anatoliens.TeilII. trionaleenuovidatisullerestantipopulazioni Senckenbergianabiologica54,227-290. Italiane. Annuario Instituto e Museo di Zool- Filippucci,M. G.; Nascetti,G.; Capanna,E.; ogiadell'UniversitadiNapoli22, 17-59. Bullini,L. (1987): Allozyme Variation and Corbet,G.B. (1978): The Mammals of the Pa- systematics of European moles of the genus laearctic Region: a Taxonomie Review. Lon- Talpa (Mammalia, Insectivora). J. Mammal- don:Brit.Mus. (Nat.Hist.). ogy68,487-499. Corbet,G.B.; Hill,J.E. (1980): AWorld List of Grulich,I. (1971): Zum Bau des Beckens (Pel- MammalianSpecies.London:Brit. Mus. (Nat. vis), einessystematisch-taxonomischenMerk- Hist.). males, bei der Unterfamilie Talpinae. Zool. Corbet,G.B.; Hill,J.E. (1986): AWorld List of Listy20, 15-28. Mammalian Species. 2nd ed. London: Brit. Grulich,I. (1982): Zur Kenntnis der Gattungen Mus. (Nat.Hist.). Scaptochirus und Parascaptor (Talpini, Mam- Corbet,G. B.; Hill,J.E. (1991): AWorld List of malia).FoliaZool.31, 1-20. Mammalian Species. 3rd ed. Oxford: Oxford Gureev,A.A. (1979): Fauna SSSR, Mlekopi- UniversityPress. tayutschie, tom 4, vyp. 2. Nasekomoyadnye Demirsoy,A. (1996): Türkiye omurgalilan. Tür- (Mammalia, Insectivora). Leningrad: Nauka. kiyeomurgalifaunasmmsistematikvebiyolo- (InRussian). jik özelliklerinin arasünlmsi ve koruma ön- Honacki,J.H.; Kinman,K.E.; Koeppel,J.W. lemlerinin saptanmasi. Memeliler. Ankara: (1982): Mammal Species of the World. Law- MeteksanA.§. (InTurkish). rence:AllenPressInc. Dogramaci,S. (1988): Geographie Variation of Hutterer,R. (1993):OrderInsectivora.In:Mam- the Mediterranean mole Talpa caeca, (Mam- mal Species of the World. ATaxonomie and malia: Insectivora)inTurkey. OndokuzMayis Geographie Reference. 2nd ed. Ed. by Üniversitesi Ziraat Fakültesi Dergisi 3, 207- D. E.WilsonandD.-A.M. Reeder.Washing- 222. (InTurkishwithEnglishsummary). ton: SmitsonianInstitutionPress.Pp.69-130. Dogramaci,S. (1989a): Taxonomy and distribu- International Commission on Zoological No- tion of moles (genus Talpa: Mammalia) in menclature (1999): International Code of Turkey. Doga TU Zooloji D. 13, 204-219. (In Zoological Nomenclature. 4th ed. London: TurkishwithEnglishsummary). The International Trust for Zoological No- Dogramaci,S. (1989b): The importance of the menclature. pelvis in distinguishing of Turkish Talpa Kefelioglu,H.;Gencoglu,S.(1996):Thetaxonomy (Mammalia: Insectivori) species. Doga TU and distribution of Talpa (Mammalia: Insecti- ZoolojiD. 13, 67-76. (InTurkishwithEnglish vora) in the Black Sea region. Turkish J. Zool. summary). 20,57-66.(InTurkishwithEnglishsummary). Ellerman,J.R.; Morrison-Scott,T. C.S. (1951): Lay,D.M. (1965): A new species of mole (genus Checklist ofPalaearctic and Indian Mammals Talpa)fromKurdistanprovince,westernIran. 1758to 1946.London:Brit.Mus. (Nat.Hist.). Fieldiana:Zoology44,227-230. Description, taxanomyand distribution ofTatpadavidiana 143 Lay,D.M. (1967): A study of the mammals of Niethammer,1; Krapp,F. (1990) Handbuch der Iran resulting from the Street expedition of Säugetiere Europas. Vol.3/1. Wiesbaden: 1962-63.Fieldiana:Zoology54,1-282. Aula-Verlag. Lemen,C.L. (1983):Theeffectivenessofmethods Osborn,D.! (1964): Notes on the moles ofTur- ofshapeanalysis.Fieldiana:Zoology15, 1-17. key.!Mammalogy45,127-129. Miller,G.S. (1940): Notes on some moles from Schwarz,E. (1948): Revision of the Old-World southeastern Asia. J. Mammalogy 21, 442- moles of the genus Talpa Linnaeus. Proc. 444. Zool.Soc. London118,36-48. Milne-Edwards,A. (1867): Observations sur Sokolov,V.E.; Tembotov,A.K. (1989): Mlekopi- quelques mammiferes du nord de la Chine. tayushchie Kavkaza: Nasekomoyadnye. Annales des Sciences Naturelles. Cinquieme Moskva:Nauka. (InRussian). serie,ZoologieetPaleontologie7,375-377. Spitzenberger,F. (1976): Ein Beitrag zur Kennt- Milne-Edwards,A. (1884): Sur la Classification nis der Vogelfauna des Cilo-Sat-Massivs (Vi- des Taupes de l'ancien continent. Comptes layet Hakäri, Türkei). Ann. Naturhist. Mus. rendus des seances de l'Academie des Wien80,315-323. Sciences99,1141-1143. Stein,G.H.W. (1960): Schädelallometrien und Misonne,X. (1959): Analyse Zoogeographique Systematik bei altweltlichen Maulwürfen des Mammiferes de lTran. Mem. Inst. Roy. (Talpidae).Mitt.Zool.Mus. Berlin36, 1-48. Sei. nat. Belgique. Deuxieme serie, Fase. 59, Stein,G.H.W. (1963): Anomalien der Zahnzahl 1-157. und ihre geographische Variabilitätbei Insec- Mitchell-Jones,A.I; Amori,G.; Bogdano- tivoren: I. Maulwurf, Talpa europaea L. Mitt. wicz,W.; Krystufek,B.; Reijnders,RJ.H.; Zool.Mus.Berlin39,223-240. Spitzenberger,F.; Stubbe,M.; Thissen, Stroganov,S.U. (1948): Sistematika krotovych J.B.M.; Vohralik,V; Zima,J. (1999): The (Talpidae). Trudy Zoologicheskogo instituta Atlas ofEuropeanMammals. London:T. and 8(2). Moskva: Akademija nauk SSSR. (In A.D.Poyser. Russian). Niethammer,! (1990a): Talpa europaea Lin- Swinhoe,R. (1870a): Catalogue ofthe mammals naeus, 1758 - Maulwurf. In: Handbuch der ofChina(southofthe RiverYangtsze) andof Säugetiere Europas. Vol.3/1. Ed. by the island ofFormosa. Proc. Scient. Meetings J.Niethammer and F.Krapp. Wiesbaden: Zool.Soc. London1870,615-653. Aula-Verlag.Pp.99-133. Swinhoe,R. (1870b): Zoological notes of ajour- Niethammer,! (1990b): Talpa occidentalis Cab- ney from Canton to Peking and Kaigan. Pro- rera, 1907 - Spanischer Maulwurf. In: Hand- ceedings ofthe scientific meetings ofthe Zo- buch derSäugetiere Europas. Vol.3/1. Ed. by ologicalsocietyofLondon1870,427-451. J.Niethammer and F.Krapp. Wiesbaden: Ziegler,A. C. (1971): Dental homologies and Aula-Verlag.Pp. 157-161. possible relationships of recent Talpidae. ! Niethammer,! (1990c): Talpa caeca Savi, 1822 - Mammalogy52,50-68. Blindmaulwurf. In: Handbuch der Säugetiere Europas. Vol.3/1. Ed. by !Niethammer and Authors' addresses: F.Krapp. Wiesbaden: Aula-Verlag. Pp. 145— Boris Krystufek, Slovenian Museum of Natural 156. History, P.O. Box 290, SI-1001 Ljubljana, Slove- Niethammer,! (1990d): Talpa stankovici V et nia (e-mail: boris.krystufek@uni-lj.si); Friederike E.Martino, 1931 - Balkan-Maulwurf. In: Spitzenberger, Naturhistorisches Museum Wien, Handbuch der Säugetiere Europas. Vol.3/1. P.O. Box 417, A-1014 Wien, Austria; Haluk Ed. by !Niethammer and F.Krapp. Wiesba- Kefelioglu, Ondokuz Mayis University, 55139 den:Aula-Verlag.Pp. 141-144. Kurupelit, Samsun,Turkey. Mamm. biol. 66 (2001) 144-154 Mammalian Biology © Urban & FischerVerlag http://www.urbanfischer.de/journals/mammbiol Zeitschriftfür Säugetierkunde Original investigation Bats in the Bavarian Alps: species composition and utilization of higher altitudes in summer ByJennnifer Holzhaider and A. Zahn ZoologischesInstitut, UniversitätMünchen, München, Germany ReceiptofMs. 10. 03. 2000 Acceptance ofMs. 20. 09. 2000 Abstract Habitat use and species composition ofbats in the higher altitudes ofthe Bavarian Alps were studied from May to September 1997. Five hundred buildings at altitudes between 800 and 1800 m above sea Level were surveyed; traces of bats or roosting animals were found in 189 ofthese. Bat occupation of buildings decreased at elevations higher than 1300 m above sea Level, and aLso decreased with increasing distance of buildings to the surrounding forest. 203 solitary roosting animals and 14 nursery colonies (indicated with*) ofthe following species were found: Myotis myotis, M. emarginatus, M. mystacinus*, M. brandti, Pipistrelluspipistreltus*, Pipistreltus nathusii, Eptesicus nilssonii*, Piecotusauritus, and Vespertib'o mun'nus. Relative to adjacent Lower regions whereMyotis myotis is the most abundant species, M. mystacinus and M. brandtitogether make up an average of 70% ofall reliably determined animals at higher altitudes. Within all occurring species, higher regions are mainly inhabited by adult males. They might thus avoid competition with nursery colonies in the lowlands. Key words: Bats, altitudinal distribution, Bavarian Alps, species composition Introduction The Bavarian Alps are likely to provide al. 1997; Güttinger 1994). However, cer- good habitats for bats in summer with ex- tain aspects in the life cycle such as repro- tensively developed pastures and large duction seem to be concentrated at lower areas of natural forests. However, lower altitudes (Spitzenberger 1993a,b), possi- temperatures and shorter Vegetation peri- bly because juvenile growth is favoured by ods relative to lowlands (Reisigl and Kel- warm conditions in many bat species (Tut- ler 1987) might be rather disadvantageous tle and Stevenson 1982; Zahn 1999). Also, for these aerial insectivores. the composition of species is influenced by Studies from neighbouring mountainous altitude (Spitzenberger 1993a; Arlettaz countries like Austria and Switzerland show et al. 1997; Güttinger 1994). This effect that higher altitudes are infact inhabitedby may depend both on climatic circumstances bats (Spitzenberger 1993a,b; Arlettaz et and on the availability of suitable roosts. 1616-5047/01/66/03-144$15.00/0. Batsinthe Bavarian Alps 145 In the "lowland areas" adjacent to the Ba- sites forbats in the lowlands, were rarelypresent varian Alps, the bat fauna is well known in the investigated buildings and are therefore (Richarz 1986; Richarz et al. 1989; Zahn notincludedinthe list ofpotentialroosts. and Krüger-Barvels 1996; Zahn and Mai- Faeces and dead or living animals were taken as er 1997). At least 10 of 16 lowland species, evidenceforsite use. Substantial accumulationof faeces was taken as evidence for colonies. Site which could potentially occur also at higher use inferred from faeces1 presence is referred to altitudes, are known to prefer various as "indirect proof", whereas alive animals were anthropogenic structures as day roosts (Ri- takenas"directproof".Whenpossible,livingani- charz 1986). In this study we therefore fo- mals were caught for species identification. With cussed on bats roosting in buildings. Build- the exception of Myotis myotis whose droppings ings exist in relatively large numbers even reliably could be identified through size and at higher altitudes in the Bavarian Alps structure, no species identificationwas attempted and are well suited for a systematic investi- using faeces. Düring the study we did not distin- guish between Pipistrellus pipistrellus and the gationofthe batfauna. Moreover,buildings "new species" Pipistrellus "pygmaeus/mediterra- are readily accessible and ofbetter compar- neus" (Häussleret al. 2000). ability than natural roosts such as rock cre- For comparison with the lowland bat population, vices or tree holes. Of special interest were data of the ASK Bavaria (database "Arten- und the following questions: (i) Which building Biotopschutzkartierung" of the Bavarian State dwelling species of bats occur over 800m Office of Environmental Protection) were called above sea level in the Bavarian Alps? (ii) upon. These data were obtained as well by sys- Does a species-specific altitudinal distribu- tematic controls of churches, Castles and other How buildingsthatoffersuitable atticsasbyroostcon- tion exist? (iii) are roosts in buildings trols conducted after house owners informed the at different altitudes used by bats? (iv) relevant authorities about bat presence. These What types ofroosts are preferred? roost controls (visits of attics, inspection of crevices) were conducted in the same way as in the present study. Material and methods The study area ranges from Garmisch-Parten- Results kirchen (11°06' O, 47°29'N) in the west to Berch- tesgaden (13°0'O, 47°37'N) in the east and in- Species composition and abundance 2cl9u6d3esmaalbtiotvuedesseafrleovmela(pZpurgosxpiitmzaet)e.lyAb8o0u0tm50t%o ofroosting bats of the area is covered with forest (mostly Fagus At 189 (41.9%) ofthe 451 potentially suita- sylvatica, mixed with Picea abies, Abies alba and ble buildings evidence of site use by bats Acer pseudoplatanus). The remaining area con- was obtained. In 100 ofthese cases, the ani- soivsetsroafboeuxtte1ns8i0v0elmyaubsoevdemsoeuantleavieln,poafstuunruessedanadl,- malaolnse'. Apretsoetnalceofw2a0s3 rionodsitciantgedanibmyalfsae(ceexs- pine meadows, dwarfpines androckyoutcrops. Five hundred buildings (from small wooden cluding nursery colonies) was found. Of mountain cabins to hotel-sized stone houses) at these, 110 were identified to the following altitudes between 800m and 1800m were sur- nine species: Greater mouse eared bat veyed between May and September 1997. Of (Myotis myotis, 4%), Geoffroy's bat these, 451 provided potential roosts for bats. As (M. emarginatus, 1%), whiskered bat potential roosts we regarded all kinds ofcrevices (M. mystacinns, 50%), Brandt's bat ifthey were dry, narrow (<5cm wide), dark and (M. brandti, 13%), common pipistrelle (Pi- without a strong airflow (<15cm deep). As de- pistrellus pipistrellus, 13%), Nathusius' pi- scribed below, these crevices were roughly di- pistrelle (Pipistrellus nathusii, 1%), north- vqiuednetdliyntuosefidvebycatbeagtosri(eRs.icShhautrtzer1s9,8w6h)iicnhsaprietefroef- ern bat (Eptesicus nilssonii, 13%), common that fact that they offer less shelter from wind long eared bat (Plecotus auritus, 5%) and andrainthanthe othertypes,were alsoincluded. parti-coloured bat (Vespertilio murinus, Large accessible attics, which are common roost 2%). With 29 animals, no reliable differen- h 146 Jennnifer Holzhaider and A. Zahn tiation between M. mystacinus and species (Fig. 2). Together they made up M. brandti could be made. We assumed that 70.5% of all reliably identified individuals. the relative fraction of the two species was Assuming a relation of 20% M. brandti in the same as that determined reliably, i.e. all M. my./br. individuals, also Brandt's bat M. mystacinus about 80%, M. brandti about was more abundant than any other species 20%. In the following, these two species are except the whiskered bat. Itis also apparent mostly combined and referred to as M. my./ that, within the studied ränge ofaltitude, no br. 64 animals remained unidentified but species showed any preference for particu- certainly belonged to smaller species (i.e. lar elevations. The only exception was larger species like M. myotis, Eptesicus M. myotis, which was found exclusively be- spec, V. murinus or Plecotus spec. could be low 1200m. However, with M. brandti and excluded). Species dwelling in buildings E. nilsonii two species were present which which occur in adjacent areas but were not are only rarely found in the adjacent low- found in buildings at higher altitudes in- land areas. cluded barbastelle (Barbastella barbastel- The highest site where roosting bats were m lus), noctule bat (Nyctalus noctula), sero- found was at 1670 in a group of cabins tine bat (Eptesicus serotinus) and lesser near Königssee (Berchtesgaden). In five horseshoe bat (Rhinolophus hipposideros). different cabins, a total of 10 animals, in- cluding M. mystacinus, PI. auritus and V. murinus was found. A Altitudinaldistribution total of 14 nursery colonies was found. Nine colonies were indicated through Bats preferred roosts in buildings at lower faeces and could thusnotbe reliablyidenti- altitudes. Figure 1 shows that the percen- fied on species level. However, concerning tage ofbuildings with traces ofbats- either pellet size they most likely belonged to Pi- alive animals or faeces - decreased signifi- pistrellus spec. or M. my./br. The other co- cantly at altitudes higher than 1300m lonies consisted of M. mystacinus (n = 2), above sea level (X2 = 26.6, p < 0.01). P. pipistrellus (n = 2) and E. nilssonii Species composition did not change signifi- (n = 1). While solitary animals reached alti- cantly with altitude. At all altitudes, tudes up to 1670m, the highest nursery col- M. my./br. were by far the most abundant ony was recognized at 1400m. Ten of the — —— —— — |— 1 i 1 | i 1 | i 1 <1000 1000-1099 1100-1199 1200-1299 1300-1399 1400-1499 >1500 n=67 n=64 n=61 n=92 n=62 n=66 n=39 Altitude[m] Fig. 1. Percentageofoccupied buildingsatdifferentaltitudes. N=numberofinvestigatedsuitablebuildings. Bats in the Bavarian Alps 147 UM. mystacinus HM. my./br. Indet. SM. brandti E. nilssonii Pipistrellus spec. PI. auritus Others 100% 90% -- o peci 80% -- w 70% -- o - 60% ü c TO 50% -- "CD 40% -- -Q (OÜ 30% -- > CO 20% -- 10% -- 0% <1000 1000-1099 1100-1199 1200-1299 1300-1499 >1500 n=37 n=20 n=20 n=24 n=24 n=15 Altitude Fig. 2. Relativeabundanceandaltitudinaldistribution ofreliablyidentified species. N=numberofbats. The altitudes 1300 m-1499 m are drawn together because of only two direct observations between 1300 m and 1399 m. Table 1: Altitudinaldistributionofnurserycolonies M. myst. E. nilssonii P. pipistrellus indicatedthrough Total faeces <1000m 1 1 5 7 1000m-1099 m 1 1 1 3 1100m-1199 m 1200m-1299 m 3 3 1300m-1399 m 1 1 Total 2 1 2 9 14 m fourteen colonies occurred below 1 100 nus were found, one consisting ofthree ani- (Tab. 1). mals (2 females, one male), the others of one male and one female each. Two sub- Age structure and composition ofsexes adults ofM. brandti (one male, one female) were found outside colony sites at the be- In all species, the great majority of solitary ginning of September. animalsconsistedofadultmales. Only 16fe- males (14.5%) were found outside nursery colonies, the first one onJuly23rd. Theybe- Preference ofroostsand surrounding longed to M. mystacinus (n = 9), M. brandti (n=2),M. myst./brandtiindet. (3),E. nilsso- The occupation ofeach type ofroost by bats nii (n= 1) andRpipistrellus (n= 1). deviates significantly from the distribution Three mating communities of M. mystaci- of existing roosting possibilities (X2-Test, 148 Jennnifer Holzhaider and A. Zahn Roof bar Roof other crevices Shutters Shingles Fig.3. Distribution ofrecordings comparedto the distribution ofpotentialroosts. Asthe numberof"othercrevices"can hardlybeestimated, novalueisgivenforthiscategory. X2 = 17.7, p<0.01, Fig. 3), which can be Concerning the compass bearings of the roughly divided into the following cate- roosts. no absolute preference for roosts, gories. The most frequently used type of openings towards one distinct orientation buildingroost (174of268recordings, includ- could be shown. However, when only ing direct and indirect evidence) was the roosts at cabins built with one distinct ori- crevice behind the board at the ridge of the entation (i.e. for example all buildings fac- roof, which was present at almost every sur- ing in North/South-direction) were consid- veyed building. It was followed in prefer- ered, southern and south-western exposed ence by crevices in the roof itself between roosts were clearly preferred to northern tin roofcovering and underlying roofbeams and north-eastern ones (X2= 4.3, p<0.05). (used by 47 animals) which are characteris- tic especially for smaller, traditionally built wooden cabins. In many buildings also Influence ofseasons Spaces behind shingles (7 records), open shutters (7 records) and other crevice-like Significantly more buildings were occupied structures (34 records) were used by bats. by bats as the summer progressed. The per- Figure 4 shows a typical mountain cabin centage of buildings with evidence of bats with the most common types of roosting rose from 7.6% (n = 13) in May to 40.1% possibilities. Also, bats significantly fa- (n = 332) in June/July, and 51.8% (n= 106) voured buildings closer to forests. While in August/September (X2 = 0.42, p<0.05). 45.6% of the roost sites in buildings less With the exception of E. nilssonii, which m than 100 away from a forest edge were was found only after the end ofJuly, the oc- occupied by bats (n = 375), only 24% currence of all species was evenly distribu- (n = 76) ofall suitable buildings farther than ted over the summer. 100m away were used (X2 = 15.4, p<0.01). Batsinthe Bavarian Alps 149 Fig.4. Typical Bavarian mountain cabin with various roosting possibilities. rb=crevice behind ridge board, rc=roofcrevice (i.e. crevices between tin covering and underlying roofbars), sh=crevice behind shutter, oc= othercrevices. Comparison between higheraltitudes is apparent between the lowlands and high- and lowlands er altitudes. In lower regions, Myotis myotis dominates Species composition at higher altitudes de- species' abundance with about 62% of all monstrates some striking differences rela- solitary animals (n = 277). Atotal of27 nur- tive to the conditions in the contiguouslow- sery colonies is known in the adjacent rural land areas. In the latter, the following districts. Also Plecotus auritus occurs quite species were found by Richarz (1986), Ri- often (17%). charz et. al (1989), Zahn andKrüger-Bar- On the otherhand, at higheraltitudesthere is vels (1996) and Zahn and Maier (1997) a strong dominance ofthe two "moustached,, (species where nursery colonies are known bat species, Brandt's bat and whiskered bat. are marked with*): Lesser horseshoe bat At higher elevations, such roost sites as (Rhinolophus hipposideros)*, greater mouse large church attics were not found. When eared bat*, Bechstein's bat (M. bech- species that normally roost in these sites in steini), Daubenton's bat (M. daubentoni)*, the lowlands are excluded (M. myotis, PI. Brandt's bat*, whiskered bat*, Geoffroy's auritus, R. hipposideros), the moustached bat*, Natterer's bat (M. nattereri)*, com- bats represent 54% of the solitary animals mon long eared bat*, barbastelle*, serotine in the mountainous study areas, but only bat, noctule bat (Nyctalus noctula), com- 22% in the lowlands (Fig. 5). P. pipistrellus, mon pipistrelle*, Nathusius' bat and parti- which reached a frequency of only 14% at coloured bat. higher altitudes, made up 28% ofall species Seven of these species were absent at in the lowlands. Also M. emarginatus and buildings in the study area, viz. R. hipposi- V. murinus are more abundant in lower deros, B. barbastellus, M. nattereri, M. dau- regions. bentoni, M. bechsteini, N. noctula and E. serotinus. E. nüssonii was found at mon- tane sites but was virtually absent in the Discussion lowland roosts. Apart from these absent species, a compari- Roosts sonbetweenthe database ASK andthepre- In general, higher altitudes in the Alps are sent study shows that a complete shiftin the inhabited by bats, although there are some relative dominance of the different species restrictions. 150 Jennnifer Holzhaider and A. Zahn m m > 800 < 800 ii ü Q n -i r t 1 r i r Q. c "U c n 2 s od £ Fig. 5. Comparison of the species composition in solitary roosting bats at high (>800m, n=101) and low (<800 m, n=50) altitudes. Af. myotis, PL auritus and R. hipposideros, which probably find only few suitabLe buildingsin hMigherregions, areexcluded. P.p.=Pipistrelluspipistrellus; P.n.=P. nathusii, M.d.=Myotisdauben- toni, M.n.= nattereri, M.e.=Af. emarginatus, B.b. =Barbastella barbastellus, V.m.= Vespertilio murinus, E.n.=Eptesicusnilsonii, N.n. =Nyctalusnoctula. The number of occupied roosts in buildings The special climatic conditions in the Alps decreases significantly over 1300m above may also be the reason for the low number sea level. This is probably due to adverse of bats roosting behind shutters, which are climatic conditions at higherelevations. The often used by whiskered bats, pipistrelles average temperature for July, for example, andbarbastelles inother areas (Spitzenber- is 17°C at 600 m but only 13°C at 1200m ger 1993a,b). Probably, their exposure to (Reisigl and Keller 1987). Furthermore, all sides affects their suitability as a roost buildings at these altitudes are often at sites in the adverse weather conditions ofhigher exposed to wind, precipitation and cold altitudes stronger than it does in lower re- temperatures, factors which might thus di- gions. However, the strong preference for minish theirsuitability as roosts. The impor- crevices behind roofbars that was observed tance ofwarmer roosts is also supported by inthis studymaypartiallybe due to ameth- the fact that, given the opportunity, bats odological artefact, since this roost type is tend to choose southern and south-western very easy to investigate. Other potential roosting possibilities at the buildings. A re- bat roosts, such as crevices in the roofitself, duced exposure to wind could also explain which offer warm shelters too, are probably the observed preference for houses in or occupied more often than is apparent, since near forests. Additionally, forests offer a bats may roost very hidden at these sites sheltered flight path between roost and and are easily overlooked. foraging area. Manybat species avoidcross- ing open land without structures such as hedges or tree lines (Limpens and Kapteyn Speciescomposition 1991). This may additionally lower the va- lue as bat roosts ofhouses located far from Some of the species absent in higher areas, forests. such as Natterer's bat, Bechstein's bat and Bats in the Bavarian Alps 151 Daubenton'sbat, are knowntoroostprefer- M. myotis, R. hipposideros, and R auritus, entially in natural structures like tree holes which were rare or absent in the higher re- (Richarz 1986), so it is not amazing that gions, depend strongly on large attics for they were not found in the surveyed build- roosting which are much more abundant in ings. However, mistnetting at several caves lower regions. Moreover, M. myotis, which in the Bavarian Alps shows that these spe- was the dominant species in the lowlands, ciesactuallydooccurathigheraltitudesalso is known to be strongly thermophilous insummer: M. bechsteiniwas caughtseveral (Spitzenberger 1988; Güttinger 1994; Ru- times at a cave near Frasdorf (1200m) dolph and Liegl 1990). In Switzerland and (Meschede pers. comra.); M. daubentoni, Austria, their complete live cycle is concen- occurredatacavenearKochelingreatnum- trated in relatively low areas. Solitary ani- m bers and could regularly be seen foraging at mals ränge up to an average of 531 in ponds up to 1100m; M. nattereri even oc- Austria, whereas nurserycolonies are found m m curred at a cave 1800 high in August and only up to 439 (Spitzenberger 1988). September 1997. Also most colonies in Switzerland occur Nyctalus noctula, which tends to roost in lower than 600m (Haffner and Moesch- trees and buildings, was neither found in ler 1995). More or less the same is true for buildings in the study area nor was it ever the lesser horseshoe bat, which mainly oc- m m seen hunting or identified acoustically in curs between 600 and 900 in Austria occasional bat-detector surveys. The ab- (Spitzenberger 1995). It is also one of the sence ofroostingnoctules in studybuildings rarest bat species in Bavaria with only two might be due to a lack ofsuitable roosts, as known nursery colonies (Zahn and these bats prefer large crevices in high Schlapp 1997). Thus, they could not be ex- buildings (Zahn et al. 1999). pected to occur in the studied buildings. Barbastella barbastellus, on the other hand, Particularly solitary animals of Plecotus is known to live at higher altitudes in Swit- auritus are also knownto use tree holes and zerland and Austria, and to preferably use bird- or bat-boxes as roosts, which were not roosts of the type that mainly occur in the investigated in this study. Theymight there- surveyed buildings: crevices behind ridge- fore be more numerous in the Alps than is bars and open shutters (Spitzenberger apparent from this investigation. 1993b). It is thus a species almost predes- The whiskered bat is the only species ofthis tined to occurinbuildings inthe study area. study forwhich every aspect ofthe life cycle Several animals caught while mistnetting at (breeding, mating. hunting, hibernating) caves near Kochel, Frasdorf, and Bichlersee was observed to take place at higher alti- in 1997 show that barbastelles actually do tudes. The distribution ofwhiskeredbat and occur in the Bavarian Alps (Meschede and Brandt's bat in other countries show that Rudolph pers. comm.). Nevertheless, theyareindeedsturdyspeciesinadverse cli- whereas it is not very numerous but widely mates: in Scandinavia they reach 64° oflati- spread in Austria (Spitzenberger 1993b), tude (Schoberand Grimmberger 1987), and it is one of the rarest species in Germany inSwitzerlandnurserycolonies ofM. mysta- m with only five recordings of nursery colo- cinus occur up to 1670 (Zingg and Bur- nies in southern Bavaria and very few re- khard 1995). In a recent investigation of cordings of solitary animals (Rudolph et the National Park Hohe Tauern in Austria al. 2001). We might have failed to find their M. my./br. also make up almost two-thirds roosts in buildings simply due to the low ofthespeciescompositionathigheraltitudes population density of this species. It is also (Hüttmeir and Reiter 1999). P.pipistrellus, possible that they use natural roosts more on the other hand, has quite similar habitat frequently thanis nowrealised, since radio- demands and roost preferences but reaches tracked individuals favoured roosts behind only61° oflatitude in Scandinavia (Schober patches of loose tree bark (Steinhauser andGrimmberger 1987) andseemstoprefer 2001). the lower altitudes. In Canton Wallis (Swit-

Bats in the Bavanan Alps: species composition and utUization of higher altitudes in summer

Similar Bats in the Bavanan Alps: species composition and utUization of higher altitudes in summer